Opening Holes in the Blanket of Inhibition: Localized Lateral Disinhibition by VIP Interneurons

Karnani, Jackson, Ayzenshtat, Hamzehei Sichani, Manoocheri, Kim, Yuste · The Journal of Neuroscience · 2016 · neuroscience · 10.1523/jneurosci.3646-15.2016 · record Karnani2016-opening-holes-in-the-blanket-of-inhibition-l
tag layer · index · concepts · authors · עברית
Where this record's text came from. faceted 2026-08 by reading the locally held text (html); every evidence string verified as an exact substring of that text before merging. Same method caveat as the companion Karnani 2016 paper: 'electrophysiology' is anchored to the slice recordings, but the headline in vivo result uses two-photon calcium imaging plus single-cell two-photon optogenetic activation, which the vocabulary cannot express. Claim type is 'mechanism' (disinhibition happens because VIPs inhibit SOMs which inhibit PCs); the quantitative ~120 um reach could equally be read as an 'existence' measurement. The 'spotlight of attention' framing is offered as speculation, not a tested claim, so it is not tagged. 2026-08-13, 2026-08-15 and 2026-08-25 schema-0.3 passes: every quote verified as an exact substring of the staged text and anchored at merge time; concept terms verified present; every edge endpoint an existing node.
6 quotations on this page contain a reference number where the paper printed a superscript. This journal style prints citations as raised numerals — parameters12,13 — and the extraction flattened them into the line, so the sentence reads parameters 12 , 13 . The digits are citation markers and not part of what the author wrote. The quotes keep them for the same reason they keep every other extraction fault: a quote is reproduced exactly or it proves nothing.
How to read this page. The article itself is not reproduced here — this paper is not cleared for redistribution, so what follows is the tag layer alone. The evidence sentences are withheld with the article. The table at the foot lists every tag the record carries; each row names the facet it belongs to, and the whose words column says whether the tag records the paper's own wording or the tagger's sentence about the paper. Every tag links to each other paper in the corpus carrying it. The machine-readable edition of the same thing is the TEI file beside this one, where the tags live in standOff, and the article is at its publisher via the link below. When we have changed our mind. Some entries in the table begin with a word in capitals and the reason for it. EVIDENCE CORRECTED — the finding still stands, but we had quoted the wrong sentence from the paper, and the quotation has been replaced with the right one. RETRACTED — the entry itself was wrong; it stays on the page, marked, with the reason, rather than being deleted. WITHDRAWN AND REPLACED — the entry was wrong, and a corrected version of it appears in the entries below. Each mark carries its own reason, in brackets, because the reasons differ from one entry to the next. An entry we have WITHDRAWN is removed from this page and kept in the record, with its reason. A line reading Audit and a date is a later re-reading of the entry against its whole sentence: DOUBTFUL means it was kept, and the reason says what is loose about it. Quotation marks inside the tagger's sentence. Where a tag is the tagger's own sentence, a phrase in quotation marks inside it is borrowed from the paper — the tagger's wording around it is not. The paper's own sentence is the evidence quote for that row; on this page it is withheld with the article, and the article is at its publisher. Nothing inside quotation marks anywhere on this page is ours: if it is quoted, it is the paper's. Whose words a concept carries. A concept entry records a term the paper itself uses, so the term, the sense given for it, and its evidence are the paper's own words, quoted where the paper fixes them. This is the opposite of a sense entry in the tag table, which is the tagger's sentence about how the paper uses a term. Where a concept entry could not be grounded in the paper's words, the entry says so on its own row.
Other fields define these differently the same word, defined another way in another discipline
standard deviation 3 fields

Tags on this paper

Every tag the record carries, each linking to every other paper in the corpus that carries it. A tag appears here whether or not its words occur in the article — the spans below are only the ones a sentence in this paper licenses. Of the 12 concept tags on this paper, none is grounded in a quoted sentence. All carry the tagger's gloss of what the term means in this paper, which is not something the article can license.
blanket of inhibition disinhibition VIP SOM PV pyramidal cells lateral inhibition two-photon optogenetics C1V1 GCaMP6s spotlight of attention spiral A VIP cell inhibits nearby SOM cells, which removes the SOM-mediated blanket inhibition on local pyramidal cells and thereby lets excitation spread through that patch. The horizontally narrow VIP axon confines the disinhibitory action to a small cylindrical volume of cortex, setting the size of the hole. During locomotion many VIPs are recruited at once, so the disinhibition from any single stimulated VIP is swamped and loses its spatial structure. somatostatin parvalbumin two-photon optogenetics visual cortex layer 2/3 interneuron locomotion vasoactive intestinal peptide neocortex axon frequency-dependent disynaptic inhibition (FDDI) holes in the blanket of inhibition locomotion epoch disinhibited cells two-photon optogenetic activation distance from spiral target change in visual responses rank sum test paired t-test paired t test t-test unpaired Student's t tests correlation Pearson's correlation z-test Z test against zero standard deviation standard error of the mean mean ± SEM Z test against zero by distance bin pairwise Pearson correlation against intersomatic distance electrophysiology

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second readingalternativesan independent re-reading of a facet, recorded whether or not it agrees
relation to other workclaim_relationshow this paper stands to another: agrees, contradicts, extends, qualifies
claim typeclaim_typethe shape of the claim: existence, mechanism, method, critique, prediction, null
concept edgeconcept_edgesan asserted relation between two concepts, with the sentence asserting it
content tagcontent_tagsa finding, definition, quantity, statistic or variable named in the paper
contributioncontributionwhat the paper claims to add
controlcontrolsan experiment run to eliminate a rival explanation
limitationlimitationswhat the paper says its evidence cannot support
mechanismmechanismshow the reported effect is said to come about
methodmethodhow the paper produced its evidence
object of studyobject_of_studywhat the paper is about — the thing it investigates
phenomenonphenomenonthe effect or regularity under study
research questionresearch_questionthe question the paper sets itself
sensesensesa distinct use of a term, quoted where the paper fixes it
transfertransfersa method or concept carried in from another field

The paper

The full text is not reproduced here. This article is not cleared for redistribution, so the page carries the tag layer only. Its evidence sentences are withheld; the grounding status remains with each tag, and the article itself is at https://doi.org/10.1523/jneurosci.3646-15.2016. The reading view exists for the papers whose licence permits it.

Every tag on this page

alternatives 3 · claim_relations 6 · claim_type 1 · concept_edges 4 · content_tags 48 · contribution 1 · controls 1 · limitations 9 · mechanisms 9 · method 1 · object_of_study 1 · phenomenon 1 · research_question 1 · senses 4 · transfers 2

Relations on this page carry a quotation for the claim. Where a relation was separately evidenced — and so tested against the vocabulary for its type — the row says so; none of 4 relations here carry that mark. The rest are the tagger’s reading of the sentence quoted beside them.

facettagfieldconf.whose words
object_of_studythe spatial extent over which a single VIP interneuron disinhibits the surrounding cortical network
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object_of_studythe paper's words
research_questionhow excitatory activity can propagate at all through a cortex covered by blanket inhibition
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research_questionthe paper's words
contributionmeasures the disinhibitory reach of a single VIP cell by four independent methods and converges on about 120 microns, giving the blanket-of-inhibition problem a quantitative answer
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contributionthe paper's words
claim_typemechanism
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claim_typethe paper's words
phenomenonlocalized lateral disinhibition by VIP interneurons
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phenomenonthe paper's words
methodelectrophysiology
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methodthe paper's words
mechanismsmechanisms
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
mechanisms[0]the tagger's words
mechanismsmechanisms
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
mechanisms[1]the tagger's words
mechanismsmechanisms
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
mechanisms[2]the tagger's words
mechanismsthe problem: SOM, PV and chandelier cells inhibit most or all pyramidal cells within their axonal arbors, which in principle should stop excitation propagating at all
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
mechanisms[3]1.0the tagger's words
mechanismsthe interneuron population looks designed to block both fast and slow excitatory activity, since PV and SOM are driven by fast and slow inputs respectively
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mechanisms[4]1.0the tagger's words
mechanismsVIP cells inhibit nearby SOM cells, and SOM cells inhibit pyramidal cells, so activating one VIP opens a local hole in the inhibitory blanket
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mechanisms[5]1.0the tagger's words
mechanismstwo-photon spiral scanning excites one targeted VIP and not its neighbours, which is what makes single-cell activation possible
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mechanisms[6]1.0the tagger's words
mechanismsthe disinhibitory effect requires concurrent sensory drive - there must be activity for the hole to admit
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mechanisms[7]1.0the tagger's words
mechanismstwo possible regimes: single VIPs opening tiny patches, or a few nearby VIPs together opening larger ones
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mechanisms[8]0.9the tagger's words
limitationsAll recordings came from layer 2/3, so the conclusions do not necessarily hold for deeper layers.
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what this limits: every conclusion in the paper, by layer: AS ALL OUR RECORDINGS WERE FROM LAYER 2/3, THEY ONLY APPLY THERE, and VIPs MAY MEDIATE QUALITATIVELY DIFFERENT EFFECTS IN THE DEEPER LAYERS
limitations[0]the tagger's words
limitationsSlice measurements of spatial reach may underestimate the true extent because processes are cut.
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what this limits: the spatial-reach measurement: OUR IN VITRO RESULTS MAY BE UNDERESTIMATES DUE TO THE INEVITABILITY OF CUT PROCESSES IN BRAIN SLICES
limitations[1]the tagger's words
limitationsThe correlational in vivo data are directionally ambiguous: nearby cells could be driving the VIPs rather than the reverse.
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what this limits: the direction of the in vivo effect: the results are consistent with VIPs disinhibiting nearby cortex BUT MIGHT EQUALLY ARISE FROM the nearby cells driving the VIPs
limitations[2]the tagger's words
limitationsall recordings are from layer 2/3 and the authors say the conclusions only apply there
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what this limits: the same scope as limitations[0] - layer 2/3 only
limitations[3]1.0the tagger's words
limitationsslice results may underestimate the reach because processes are cut
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what this limits: the same scope as limitations[1] - cut processes underestimate reach
limitations[4]1.0the tagger's words
limitationsthe correlational results could equally reflect nearby cells driving the VIPs rather than the reverse, which is why the optogenetic experiment was needed
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duplicate of: limitations[2] · what this limits: the same scope as limitations[2] - the correlation does not fix the direction
limitations[5]1.0the tagger's words
limitationsduring locomotion the spatial structure disappears, attributed to many more VIPs being recruited
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what this limits: the spatial-structure result outside quiescence: DURING LOCOMOTION EPOCHS THE DISTANCE DEPENDENCE WAS SCRAMBLED, attributed to recruitment of many more VIPs
limitations[6]1.0the tagger's words
limitationssix cells were inhibited rather than disinhibited and were set aside as under 1% of the sample
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what this limits: the completeness of the disinhibition account: BECAUSE THE INHIBITED CELLS FORMED <1% OF ALL CELLS, WE DID NOT ANALYZE THEM FURTHER, THOUGH WE NOTE THEY MAY BE SOMS - cells that behaved oppositely were set aside
limitations[7]1.0the tagger's words
limitationsthe requirement for sensory stimulation is supported by data not shown
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what this limits: the stated requirement for sensory stimulation: it WAS REQUIRED FOR SEEING THE DISINHIBITORY EFFECT OF SINGLE VIP ACTIVATION (DATA NOT SHOWN) - the condition for the central effect rests on data not in the paper
limitations[8]0.9the tagger's words
transfersthe spotlight metaphor, imported as the functional interpretation of a local disinhibitory hole
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transfers[0]1.0the tagger's words
transfersattractors, offered as what selectively sculpted circuits might implement
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transfers[1]0.85the tagger's words
controlsthe same analysis re-run triggering on the other cells, mostly pyramidal cells, instead of the original trigger cells
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eliminates: a result that follows from which cell was chosen as the trigger rather than from the population structure · outcome (stated by the tagger, no quotation): held
controls[0]0.85the tagger's words
sensesblanket of inhibition is the specific claim that SOM, PV and chandelier cells inhibit most or all pyramidal cells within their arbors indiscriminately - it is what makes disinhibition necessary rather than merely available
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senses[0]1.0the tagger's words
sensesthe hole is measured four different ways - axon extent, paired connectivity, activity correlation, and single-cell optogenetics - and reported as a radius of about 120 microns, equivalently a diameter of about 240
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senses[1]1.0the tagger's words
sensesspotlight of attention is used as an analogy for what a local disinhibitory hole could implement, not as a claim that it does
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senses[2]1.0the tagger's words
sensesdisinhibited is defined operationally as a visual response increasing by more than two standard deviations during VIP activation
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senses[3]1.0the tagger's words
alternativesclaim_type: mechanism
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a second reading reaching the same value. Recorded: mechanism — note: a second reading reaching the same value from a different sentence. Not applied - a person decides.
alternatives.claim_typethe tagger's words
alternativesmethod: two-photon optogenetic activation of individual VIP cells with C1V1 during calcium imaging in vivo, plus paired slice recordings and morphological reconstruction
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a second reading DIFFERING from the recorded value. Recorded: electrophysiology — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides.
alternatives.methodthe tagger's words
alternativesphenomenon: lateral disinhibition in neocortex
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a second reading DIFFERING from the recorded value. Recorded: localized lateral disinhibition by VIP interneurons — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides.
alternatives.phenomenonthe tagger's words
claim_relationsextends → Kapfer et al. (2007), Silberberg and Markram (2007) and Murayama et al. (2009) on SOM-mediated frequency-dependent disynaptic inhibition
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claim_relations[0]the tagger's words
claim_relationsreplicates → Zhang et al. (2014) on the uniquely localized disinhibitory effect of VIP cells
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claim_relations[1]the tagger's words
claim_relationsextends → Pi et al. 2013 and Fu et al. 2014 on the VIP to SOM disinhibitory motif - both independently tagged in this corpus
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claim_relations[2]1.0the tagger's words
claim_relationssupports → Zhang et al., who also found VIPs uniquely localised with a radius under 200 microns
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claim_relations[3]1.0the tagger's words
claim_relationssupports → morphological reports that VIP axons are horizontally restricted
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claim_relations[4]0.95the tagger's words
claim_relationsextends → Fu et al. 2014 on locomotion driving VIP activity, by showing the spatial structure is lost in that state
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
claim_relations[5]1.0the tagger's words
concept_edgesVIP —causes→ SOM
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
strength of the relation: 0.95 · note: inhibits. same edge as Fu2014 and Pi2013 - three corpus papers now carry it, from three labs
also in: VIP · SOM
concept_edges[0]1.0the tagger's words
concept_edgesSOM —causes→ pyramidal cells
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
strength of the relation: 0.95
also in: SOM · pyramidal cells
concept_edges[1]1.0the tagger's words
concept_edgesVIP —operationalised-as→ spotlight of attention
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
strength of the relation: 0.4 — the record marks this weak · note: explicitly an analogy - may therefore enable. the low weight carries that this is offered, not shown Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but VIP interneurons 'MAY' enable propagation by generating a spotlight of attention; operationalised-as is loose and hedged.
concept_edges[4]1.0the tagger's words
concept_edgesSOM —operationalised-as→ lateral inhibition
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status: ATTRIBUTED, NOT MEASURED HERE: the span reports what the field holds - 'disynaptic lateral inhibition between nearby pyramidal cells' 'Quotation withheld' - with three citations (Kapfer et al. 2007; Silberberg and Markram 2007; Adesnik et al. 2012). This paper's own title is 'Opening holes in the blanket of inhibition', so the received account is the thing it goes on to complicate. · strength of the relation: 0.9 · note: RE-GROUNDS another row of this record: the old span named ONE endpoint. Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but lateral inhibition 'IS THOUGHT TO work through' SOMs - a mechanism; operationalised-as is loose.
concept_edges[5]0.9the tagger's words
content_tagsrank sum test
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also in: rank sum test
content_tags.statistics[0]the tagger's words
content_tagspaired t-test
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also in: paired t-test
content_tags.statistics[1]the tagger's words
content_tagst-test
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: t-test
content_tags.statistics[2]the tagger's words
content_tagscorrelation
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: correlation
content_tags.statistics[3]the tagger's words
content_tagsz-test
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: z-test
content_tags.statistics[4]the tagger's words
content_tagsstandard deviation
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.statistics[5]the tagger's words
content_tagsstandard error of the mean
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.statistics[6]the tagger's words
content_tagsrank sum test
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: rank sum test
content_tags.statistics[7]1.0the tagger's words
content_tagsZ test against zero by distance bin
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.statistics[8]1.0the tagger's words
content_tagspairwise Pearson correlation against intersomatic distance
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content_tags.statistics[9]0.9the tagger's words
content_tagsOptogenetically silencing SOM interneurons abolished the disynaptic inhibition between paired pyramidal cells, giving direct loss-of-function evidence that SOMs mediate lateral inhibition.
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content_tags.findings[0]the tagger's words
content_tagsActivating VIP interneurons during the presynaptic spike train also blocked the disynaptic inhibition, showing VIPs can veto lateral inhibition.
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content_tags.findings[1]the tagger's words
content_tagsReconstructed VIP axons were narrower than SOM or PV axons and descended away from the pia.
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content_tags.findings[2]the tagger's words
content_tagsSOMs postsynaptic to a VIP were significantly closer to it than unconnected SOMs, so VIP inhibitory output is spatially restricted.
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content_tags.findings[3]the tagger's words
content_tagsTwo-photon activation of a single VIP in vivo disinhibited visual responses selectively in cells near the stimulated VIP.
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content_tags.findings[4]the tagger's words
content_tagsThe disinhibitory reach of one VIP cell measured about 120 μm in radius, i.e. a hole roughly 240 μm across.
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content_tags.findings[5]the tagger's words
content_tagsSpontaneous VIP activity was closely tied to activity within a 50 um radius, whereas pyramidal-cell activity showed only weak distance dependence.
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content_tags.findings[6]the tagger's words
content_tagsThe spatial localization of the single-VIP effect disappeared during locomotion.
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content_tags.findings[7]the tagger's words
content_tagsdisinhibited cells sat significantly closer to the activated VIP than cells in general, 66 against 101 microns
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content_tags.findings[8]1.0the tagger's words
content_tagsthe effective disinhibitory radius of a single VIP is about 120 microns
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content_tags.findings[9]1.0the tagger's words
content_tagsequivalently, VIPs open holes of about 240 microns diameter in the SOM blanket
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content_tags.findings[10]1.0the tagger's words
content_tagsconnected VIP-SOM pairs sat closer together than unconnected ones, 112 against 163 microns
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content_tags.findings[11]1.0the tagger's words
content_tags33 of 638 cells were disinhibited and 6 were inhibited by single-VIP activation
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content_tags.findings[12]1.0the tagger's words
content_tagsthe spatial structure of the effect disappears during locomotion
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content_tags.findings[13]1.0the tagger's words
content_tagstargeting a neighbouring VIP did not excite the recorded one, establishing single-cell resolution
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content_tags.findings[14]1.0the tagger's words
content_tagsblanket of inhibition
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.definitions[0]the tagger's words
content_tagsdisinhibition
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: disinhibition
content_tags.definitions[1]the tagger's words
content_tagsfrequency-dependent disynaptic inhibition (FDDI)
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.definitions[2]the tagger's words
content_tagsholes in the blanket of inhibition
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content_tags.definitions[3]the tagger's words
content_tagslocomotion epoch
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content_tags.definitions[4]the tagger's words
content_tagsblanket of inhibition
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.definitions[5]1.0the tagger's words
content_tagsdisinhibited cells
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content_tags.definitions[6]1.0the tagger's words
content_tagstwo-photon optogenetic activation
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content_tags.variables[0]the tagger's words
content_tagsdistance from spiral target
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content_tags.variables[1]the tagger's words
content_tagschange in visual responses
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content_tags.variables[2]the tagger's words
content_tags33/638 cells — cells whose visual responses were significantly disinhibited by single-VIP activation, out of all imaged cells
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content_tags.quantities[0]the tagger's words
content_tags∼240 μm — diameter of the disinhibitory hole a VIP opens in SOM-mediated blanket inhibition
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[1]the tagger's words
content_tags440 ± 53 μm, 233 ± 29 μm — vertical and horizontal axon extent of VIP cells, against 756 ± 85 / 615 ± 82 μm for SOMs
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content_tags.quantities[2]the tagger's words
content_tags41 ± 16 ms — onset latency of the disynaptic inhibition in the second pyramidal cell after the spike train began
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content_tags.quantities[3]the tagger's words
content_tagssix of eight — individual VIP cells successfully driven to spike by targeted two-photon spiral stimulation in slices
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content_tags.quantities[4]the tagger's words
content_tags112 ± 25 μm vs 163 ± 14 μm, p = 0.01 — intersomatic distance of SOMs connected versus not connected to a recorded VIP
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content_tags.quantities[5]the tagger's words
content_tags∼233 μm/2 = 116 μm ... ∼112 μm — convergent anatomical and functional estimates of a single VIP's lateral reach
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[6]the tagger's words
content_tagsabout 120 microns — effective disinhibitory RADIUS of a single VIP cell
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[7]1.0the tagger's words
content_tagsabout 240 microns — the same result expressed as a DIAMETER - not a discrepancy with the 120 figure
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[8]1.0the tagger's words
content_tagsaxons about 233/2 = 116 microns laterally; SOMs inhibited within about 112; correlations raised at 75 to 150; optogenetic disinhibition within 60 to 120 — four independent estimates of one quantity, converging on about 120 microns
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[9]1.0the tagger's words
content_tagsconnected 112 plus or minus 25 microns, not connected 163 plus or minus 14, p = 0.01 — VIP to SOM connection probability against distance
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[10]1.0the tagger's words
content_tagsdisinhibited 66 plus or minus 9 microns against all cells 101 plus or minus 2, p = 0.002 — how much closer the affected cells were
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[11]1.0the tagger's words
content_tags33 disinhibited, 6 inhibited, 599 unchanged of 638 cells across 27 experiments — the classification outcome
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[12]1.0the tagger's words

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© 2026 Shir Sivroni · ORCID 0009-0000-2597-2824